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Further Notes on the Port Kennedy Cave Arvicolid Rodents

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Further Notes on the Port Kennedy Cave Arvicolid Rodents Paludicola 9(1):13-20 November 2012 © by the Rochester Institute of Vertebrate Paleontology FURTHER NOTES ON THE PORT KENNEDY CAVE ARVICOLID RODENTS Robert A. Martin Department of Biological Sciences, Murray State University, Murray, KY 42071 ABSTRACT Cope (1871) named five Microtus-like species from the Port Kennedy Cave deposit of Montgomery County, Pennsylvania, all of which he referred to the genus Arvicola. Using Cope’s illustrations and descriptions, a review by C. W. Hibbard and examination of three Port Kennedy specimens, all of Cope’s species (Arvicola speothen, A. tetradelta. A dideltus, A. sigmodus and A. involutus) are deemed nomina dubia. Two available specimens are clearly referable to Pitymys cumberlandensis van der Meulen, and another is referred to cf. Microtus (Pedomys) llanensis Hibbard. On the basis of these specimens and other considerations, Port Kennedy is considered to be younger than the Cumberland Cave local fauna of Maryland and approximately contemporaneous with the Trout Cave 2 local fauna of West Virginia, falling within Great Plains Rodent Zone (RZ) 14, between about 1.3 - 0.64 Ma. The origin of Pitymys is obscure. Absence of a suitable North American ancestor and dental features of P. cumberlandensis suggest we should look for an Old World immigrant ancestor with undifferentiated enamel, a complex m3, and an m1 with five closed triangles and deep, provergent reentrants. Confluent T1-2 on lower molars and T2-3 on M3 in P. cumberlandensis may preclude this species as an ancestor for later North American pine voles. INTRODUCTION Hibbard (1955) valiantly attempted to make sense of the material from Cope’s (1871) descriptions and Six species of arvicolid rodents were described by comparisons made by Gidley and Gazin (1938) when Cope (1871) from the Port Kennedy Cave deposit in they described arvicolid material from the Cumberland Montgomery County, Pennsylvania. All were referred Cave deposit of Maryland. Van der Meulen (1978) to the genus Arvicola. The taxonomic history of these later restudied the arvicolids from the Cumberland species was reviewed by Hibbard (1955), and he Cave deposit of Maryland, referring them to Microtus synonymized five of the Microtus-like species under paroperarius Hibbard (1944) and the new species Microtus speothen, “Pedomys or Pitymys “ dideltus Pedomys guildayi and Pitymys cumberlandensis. In his and “Pedomys or Pitymys” involutus. The sixth paper, van der Meulen (1978) considered “Pitymys” Hibbard identified as an extinct muskrat, Ondatra involutus as a nomen dubium, but did not consider hiatidens, which is probably juvenile O. annectens Cope’s (1871) other species. From Hibbard’s (1955) (Martin et al., 2009). He also added the genus Neofiber descriptions and illustrations of the Port Kennedy to the arvicolid community. Hibbard examined material material, van der Meulen’s (1978) classic work on the in the American Museum of Natural History (AMNH) Cumberland Cave arvicolids and subsequent research and a few specimens from the Academy of Natural on arvicolids from northeastern states (Pfaff, 1990) it Sciences of Philadelphia (ANSP), as the Port Kennedy seems likely that the Port Kennedy Cave arvicolid specimens were split between the two institutions. The cadre includes a species of Microtus near Hibbard’s holotypes were kept at the AMNH and, according to (1944) M. paroperarius and two additional species Hibbard (1955) were in the following condition when near van der Meulen’s (1978) Pitymys cumber- he examined them: 1) Arvicola speothen (AMNH landensis and either van der Meulen’s (1978) Pedomys 8689) – imprint of a left ramus with only m3 guildayi or Hibbard’s (1944) Pedomys llanensis. The preserved; 2) Arvicola tetradelta (AMNH 8692) – first question that arises is: Are any of Cope’s according to Cope (1871) is a maxillary piece with M2- holotypes or topotypes sufficiently preserved to be M3 but Hibbard (1955) identified the molars as m2- diagnostic at the species level, perhaps supplanting one m3; 3) Arvicola involuta (AMNH 8699a) – teeth or more of Hibbard’s and van der Meulen’s later absent, presumably disintegrated. 4) Arvicola sigmodus names? Combining information from Cope’s (1871) (AMNH 8696) – molars missing, only piece of ramus original work, Hibbard’s (1955) review and personal and lower incisor remains; 5) Arvicola didelta (AMNH obervation of three specimens, I review the Port 8694) – all teeth missing, presumably disintregrated. Kennedy Microtus-like voles and tentatively correlate the Port Kennedy Cave assemblage with other early 13 14 MARTIN—PORT KENNEDY ARVICOLID RODENTS Pleistocene assemblages from the eastern and the extinct Allophaiomys, are generally indistin- midwestern states. guishable, and as such it would not be possible (at least currently) to distinguish M. paroperarius from P. METHODS guildayi based on this molar. Relying on Cope’s (1871) description of the m1-m2 of AMNH 8689 before it was Upper molars are abbreviated by upper case lost, Hibbard (1955, p. 90) further referred the letters (e.g., M3), lowers by lower case letters (e.g., ambiguous AMNH 8692, the holotype of Cope’s m3). Three arvicolid specimens from the Port Kennedy Arvicola tetradelta, to M. speothen. However, even if Cave deposit housed at the Philadelphia Academy of the two molars of AMNH 8692 are eventually Natural Sciences (ANSP) were examined and identified correctly, they cannot be used to identify the measured for this study: ANSP 141 (figured by subgenus Microtus, let alone M. speothen or M. Hibbard, 1955; fig. 2D): part right (R) mandible with paroperarius. In the absence of additonal material that incisor and m1-m3; ANSP 142 (figured by Hibbard, could be construed to be part of a “type series,” I 1955; fig. 2B): part left (L) mandible with m1; ANSP recommend we consider Arvicola speothen and A. 18907: part L mandible with m1-m3. tetradelta as nomena dubia. Measurements were made with an AO filar ____________________________________________ micrometer coupled to an AO binocular microscope. The micrometer was calibrated with a 2.0 mm AO slide and measurements were multiplied by the appropriate correction factor (0.615 in this case). Measurements made on m1 are shown in Figure 1. Ratios calculated from the measurements are as follows: A/L = 100a/L, a measure of the relative length of the ACC; W’/W = 100w’/W, a measure of the relative width of the anterior cap; B/W = 100b/W, a measure of the degree of confluency between the anterior cap and the triangles 4 and 5; C/W = 100c/W, a measure of the degree of confluency between triangles 4 and 5 (Martin, 1991). I follow van der Meulen (1978) in assuming that T1 is not expressed on M2-M3; consequently the triangles begin on these teeth after the anterior loop with T2 instead of T1 as illustrated by Pfaff (1990; fig. 3). Dental abbreviations in the text are as follows: BRA = buccal re-entrant angle, LRA = lingual re- entrant angle. Figure 2 was traced from a digital photograph, reduced and darkened. Parts of some of the triangle edges were obscured by thick glue and are therefore best estimates. Nevertheless, most triangles clearly displayed undifferentiated enamel. SYSTEMATIC PALEONTOLOGY FIGURE 1. Topography of a Microtus left m1 (after Pfaff 1990). ac FAMILY ARVICOLIDAE = anterior cap (sometimes also referred to as the acd, or anteroconid); COPE’S SPECIES L = occlusal length; a = length of ACC (anteroconid complex; all structures anterior to T3); W = greatest width ACC; w’= greatest Arvicola speothen Cope, 1871 width of anterior cap; b = width of dentine isthmus connecting T4- Arvicola tetradelta Cope, 1871 T5 and the anterior cap; c = width of dentine isthmus connecting T4 - T5; T1, T2 = triangles 1 and 2 (to identify other triangles, continue Comments—Despite the presence only of m3 counting from T2; buds from the ac are considered “incipient” triangles). (m1 and m2 are missing), Hibbard (1955) retained _______________________________________________________ AMNH 8689 as the holotype of Microtus speothen and, without explanation, suggested that M. speothen and Arvicola involuta Cope, 1871 M. paroperarius from the Cudahy and Subrite local faunas of Kansas (Hibbard, 1944) might be Comments—Molars of the holotype of Arvicola conspecific. However, the m3 of most Microtus, involuta (AMNH 8694) have disintegrated and the including the extant subgenera Microtus, Pedomys and illustration of an m1 provided by Cope (1871, fig. 16) PALUDICOLA, VOL. 9, NO. 1, 2012 15 is not definitive of any Microtus-like subgenus or 15, suggest didelta is a member of the taxon Pedomys species, and it does not resemble the m1 of extant (Martin, 1991; Pfaff, 1990). However, both the series Pitymys pinetorum, despite Cope’s suggestion to the of isolated m1s and the holotype of A. didelta are contrary (Cope, 1871, p. 90). I recommend we consider missing. The topotype m1-m3 illustrated by Hibbard Arvicola involuta as a nomen dubium. (1955) is not advanced enough in dental anatomy to be ___________________________________________ distinguishable from Great Plains Pleistocene Microtus referred to M. (Allophaiomys) pliocaenicus (Martin, 1989). Although the molars of AMNH 8688 display positive enamel differentiation (unlike Pitymys cumberlandensis, in which the enamel is undifferentiated) both M. pliocaenicus and M. (Pedomys) guildayi from Cumberland Cave display positive enamel differentiation (van der Meulen, 1978; Martin, 1989, 1995). Because the remaining material of Arvicola didelta is not sufficient to define a distinct Microtus species, I recommend that A. didelta be considered a nomen dubium. Arvicola sigmodus Cope, 1871 Comments—The molars from the holotype mandible of A. sigmodus, AMNH 8606, are missing, and none of the uncatalogued individual molars illustrated by Cope (1871, fig. 17) are currently recognized in the AMNH collections (R. Oleary, personal commmunication). Cope’s (1871) diagnosis of A. sigmodus is contradictory, indicating some similarity to species we currently recognize as Microtus (Pedomys) ochrogaster and Pitymys pinetorum, which is not surprising because he illustrated a combination of species under A.
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